Clupeiformes Meristic Characters in Species Belonging to the Order Clupeiformes Whose Adults Or Larvae Have Been Collected in the Study Area

Home , Alewife (fish), Anchoa, Anchoa hepsetus, Anchoa mitchilli, Atlantic thread herring, Clupea

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Clupeiformes Meristic characters in species belonging to the order Clupeiformes whose adults or larvae have been collected in the study area. Families and genera are listed alphabetically. Rarely observed counts in parentheses. Sources: Munroe (2000; 2002a; 2002b). Also see species accounts.

Family Dorsal Anal Pectoral Pelvic Species Vertebrae Fin Rays Fin Rays Fin Rays Fin Rays Clupeidae Alosa aestivalis 47–53 15–20 15–21 14–18 9–11 Alosa mediocris 53–55 15–20 19–23 15–16 9 Alosa pseudoharengus 46–50 12–18 15–20 14–16 10 Alosa sapidissima 55–57 (14)18–19(21) (18)21–22(25) 13–18 8–10 Brevoortia tyrannus 45–50 18–24 18–24 13–19 7 Clupea harengus 52–62 (16)17–19(22) (15)17–19(21) 13–21 6–10 Dorosoma cepedianum 47–51 10–15 25–37 14–17 7–10 Dorosoma petenense 43–44 11–15 17–27 12–17 (7)8 Etrumeus teres 46–50 15–22 10–13 14–16 8 Harengula jaguana 41–43 17–19 17–18 13–15 (7)8 Opisthonema oglinum 45–49 17–22 20–25 15–19 8–9 Sardinella aurita 45–49 16–19 16–20 15–16 7–8 Engraulidae Anchoa hepsetus 40–44 13–16 19–23 13–15 7 Anchoa mitchilli 38–44 13–17 23–30 11–12 7 Engraulis eurystole 43–45 13–16 15–18 14–16 7

Note: Three species of engraulids can best be distinguished by the relative positions of their dorsal and anal fins and by anal fin ray counts. These characters are most reliable in larvae >10 mm, juveniles and adults. Dorsal and anal fin origins can be almost opposite each other in adult A. mitchilli. Early Stages of Fishes in the Western North Atlantic Ocean 185

Clupeiformes Synopsis of characters for distinguishing larvae of the Clupeidae. "Dors-Anal" refers to number of myomeres between posterior dorsal and anterior anal fins. This number decreases at transformation in most species, as fin positions shift. Note that this number remains relatively stable through ontogeny in the genus Alosa.

Notochord Notochord Total Preanal Predorsal Dors-Anal Pigment Pigment Myomeres Myomeres Myomeres Myomeres Dorsal Tip Ventral Tip Miscellanous

Alosa aestivalis 47–53 42–45 – 11–13 No Yes Slim-bodied, small-eyed Alosa mediocris 53–55 38–42 – 7–8 In small larvae Yes – Alosa pseudoharengus 46–50 39–43 – 7–9 Yes Yes Deep-bodied, large-eyed Alosa sapidissima 55–57 41–47 – 9 to 71 No Yes – Brevoortia tyrannus 45–50 38–40, 30–31, 5 to 1 In small larvae Yes High D and A then 35–36 then 24–25 fin ray counts Clupea harengus 52–62 47, then 33 to 25 8–9 to 4 Varies Yes Late anal fin 41–46 ray formation Dorosoma cepedianum 47–51 39–44 – 10 to 71 No No Very high anal fin ray count Dorosoma petenense 43–44 36 – 10 to 71 No No Scattered pigment on caudal peduncle Etrumeus teres 48–50 39–40, 27–32, 5 to 2 No Yes Early teeth; low then 36 then 26 anal fin ray count Harengula jaguana 39–42 35, 25, 5 to 7 Yes If yes, Low myomere then 27 then 10 forms late count Opisthonema oglinum 45–49 40–41, 26–27, 8–10 to 5–7 No Yes High anal fin then 34–36 then 22–23 ray count Sardinella aurita 45–48 38–40, 26–27, 7–8 to 5 No Yes – then 34 then 18

1 Determined from illustrations; need confirmation

Patterns of ventral pigment, posterior to cleithrum, in larvae of four species of Alosa. These patterns may be incomplete in many larvae. (Walsh et al., 2005, modified from Sismour, 1994.)

Walsh et al., (2005) also provide a size- dependent key to larvae of these four species (plus Dorosoma cepedianum) from the southern part of the study area (Roanoke River, North Carolina).

A. pseudoharengus A. aestivalis A. mediocris A. sapidissima 186

Alosa aestivalis (Mitchill, 1815) Clupeidae Blueback herring

Range: Atlantic coast of North America from Nova Scotia to St. John's River, Florida Meristic Characters Habitat: Pelagic, schooling species, inhabiting coastal areas over the inner Myomeres: 47–53 continental shelf (deeper during winter), moving into brackish or fresh Vertebrae: 14–16+33–35 waters to spawn Dorsal fin rays: 15–20 Spawning: Adults migrate into coastal rivers during spring for spawning; begins in Anal fin rays: 15–21 late Apr, continues through May in southern part of range, as late as Sep Pectoral fin rays: 14–18 in Connecticut Pelvic fin rays: 9–11 Caudal fin rays: 7–8+10+9+6–7 Eggs: – Pelagic to semi-demersal, slightly adhesive – Diameter: 0.87–1.11 mm – Chorion: yellowish, semitransparent – Chorion; somewhat thick, inner surface corrugated – Oil globules small, unequal in size, scattered – Yolk: granular – Perivitelline space: wide Yolk-sac larva Larvae: – Hatching occurs at 3.1–5.0 mmTL; eye pigmented – Body elongate with long, straight gut; anus always posterior to dorsal fin

– Sequence of fin ray formation: D–C–A, 1P , P2 (ossification sequence based on illustrations) – 11–13 myomeres between posterior dorsal and anterior anal fins – Discrete melanophores below pectoral fin base become a double line of pigment – Pigment along upper and lower gut in distinct rows – Notochord tip pigment present on ventral side only – Transformation occurs between 20 and 30 mm

Note: 1. Slimmer and smaller-eyed than A. pseudoharengus 2. See comparative table in Clupeidae Introduction

Early Juvenile:

E. 45.0 mmTL

Figures: Adult: H.L. Todd (Hildebrand, 1963a); Egg and yolk-sac larva: Mansueti and Hardy, 1967; A–B: Chambers et al., 1976; C: Ann S. Green (Hildebrand, 1963a; D: Nancy D. Patton (Hildebrand, 1963a); E: Mansueti and Hardy, 1967 References: Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 187

Alosa aestivalis

A. 12.0 mmTL

B. 14.8 mmTL

C. 20.5 mmTL

D. 25.0 mmTL 188

Alosa mediocris (Mitchill, 1814) Clupeidae Hickory shad

Range: Atlantic coast of North America from Bay of Fundy to Florida Habitat: Inner continental shelf, estuaries, tidal rivers; oceanic distribution during Meristic Characters winter not well known Myomeres: 53–55 Spawning: In tidal freshwater, late Apr through early Jun Vertebrae: 17+37 Dorsal fin rays: 15–20 Eggs: – Spherical, semi-demersal, slightly adhesive Anal fin rays: 19–23 – Diameter: 0.96–1.65 mm Pectoral fin rays: 15–16 – Chorion: transparent, thick, finely corrugated Pelvic fin rays: 9 – Yolk: densely segmented, amber Caudal fin rays: 9+10+9+7 – Oil globules: few, small – Perivitelline space: very wide Larvae: – Hatching occurs at 5.2–6.5 mm; eye pigmented – Body elongate with long, straight gut; anus always posterior to dorsal fin – Preanus length 80–86% TL Yolk-sac larva – Posterior gut with muscle-band striations – Flexion occurs at 10–15 mm

– Sequence of fin ray formation: D–C, A, P2–P1 (ossification sequence based on illustrations) – Pigmentation includes elongate melanophores along ventral surface of gut with few spots on dorsal surface in early larvae; later larvae have melanophores on ventrum from throat area to region above anal fin – Transformation occurs between 20 and 30 mm Note: 1. See comparative table in Clupeidae Introduction Early Juvenile:

E. 35.2 mmTL

Figures: Adult: H.L. Todd (Hildebrand, 1963a); Egg, yolk-sac larva and A–E: Mansueti, 1962 References: Jones et al. 1978; Able and Fahay, 1998; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 189

Alosa mediocris

A. 15.8 mmTL

B. 19.5 mmTL

C. 24.3 mmTL

D. 27.0 mmTL 190

Alosa pseudoharengus (Wilson, 1811) Clupeidae Alewife

Range: Atlantic coast of North America from Newfoundland to South Carolina, most abundant between Gulf of Maine and Chesapeake Bay; also land- locked populations, Great Lakes and New York Meristic Characters Myomeres: 46–50 Habitat: Pelagic, schooling species, inhabiting coastal areas adjacent to fresh- Vertebrae: 46–50 water nursery areas; spends fall and winter in continental shelf waters Dorsal fin rays: 12–18 56–110 m deep, primarily off Southern New England, Georges Bank Anal fin rays: 15–20 and Gulf of Maine Pectoral fin rays: 14–16 Spawning: Adults migrate into coastal rivers during spring for spawning; larvae Pelvic fin rays: 9–10 and juveniles occur in estuaries throughout range, except those with Caudal fin rays: 10+9 (PrC) limited freshwater input. Eggs: – Demersal and adhesive – Diameter: 0.87–1.11 mm – Chorion: semitransparent and yellowish – Yolk: segmented – Oil globules: small, unequal in size, scattered – Perivitelline space: wide Larvae: – Hatching occurs at 3.1–5.0 mm; eyes pigmented Yolk-sac larva – Body elongate with long, straight gut; anus always posterior to dorsal fin – Preanus length about 80% TL – Posterior gut with muscle-band striations at sizes >10.0 mm – Flexion occurs at about 9.0 mmSL

– Sequence of fin ray formation: D, C – A – 2P – P1 (ossification sequence based on illustrations) – Pigmentation includes row of spots on dorsal half of anterior gut, ventral surfaces of posterior gut; early larvae have rows of spots on dorsal and ventral surfaces of caudal peduncle; 1 large melanophore above base of pectoral fin; dorsolateral pigment appears in larger larvae – Transformation occurs between 25 and 30 mm Note: 1. Deeper-bodied and larger-eyed than Alosa aestivalis 2. See comparative table in Clupeidae Introduction Early Juvenile: – Larger juveniles have single, dark spot on side posterior to opercle – Sharp scutes form along midline of belly

E. 42.0 mmTL

Figures: Adult: H.L. Todd (Hildebrand, 1963a); Egg, yolk-sac larva: Mansueti and Hardy, 1967; A–B: Chambers et al., 1976; C: Norden, 1967; D: Ann S. Green (Hildebrand, 1963); E: Mansueti and Hardy, 1967 References: Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 191

Alosa pseudoharengus

A. 9.0 mmTL

B. 15.0 mmTL

C. 24.5 mmTL

D. 29.0 mmTL 192

Alosa sapidissima (Wilson, 1811) Clupeidae American shad

Range: Atlantic coast of North America from Labrador to Florida; most abundant between Connecticut and North Carolina Meristic Characters Habitat: Congregate in Gulf of Maine during summer and fall, then most overwin- Myomeres: 55–57 ter off Middle Atlantic Bight, at depths to 230 m Vertebrae: 55–57 Spawning: Adults migrate into coastal rivers to spawn during spring; larvae develop Dorsal fin rays: 14–21 in freshwater, juveniles form schools as they drift downstream summer and Anal fin rays: 18–25 fall Pectoral fin rays: 13–18 Pelvic fin rays: 8–10 Eggs: – Demersal, adhesive early stages Caudal fin rays: 10+9 (PrC) – Later stages non-adhesive – Diameter: 2.5–3.8 mm – Chorion: Transparent, pale pink or amber – Yolk: segmented – Oil globules: none – Perivitelline space: wide Yolk-sac Larva Larvae: – Hatching occurs at 5.7–10.0 mm – Body elongate with long, straight gut; anus always posterior to dorsal fin – Air bladder obvious by 14 mm TL – Preanal finfold retained throughout larval development

– Sequence of fin ray formation: D, C – A – P2 – P1 (ossification sequence based on illustrations) – Transformation occurs between 25 and 30 mm Note: 1. See comparative table in Clupeidae Introduction 2. Ventral pigment in small A. sapidissima (center) differs from that in A. aestivalis (left) and A. pseudoharengus (right) in sizes <13.0 mmTL (after Sismour, 1994) Early Juvenile:

E. 51.8 mmTL

Figures: Adult: H.L. Todd (Hildebrand, 1963a); Egg, yolk-sac larva and A: Ryder, 1887; B–E: Mansueti and Hardy, 1967 References: Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 193

Alosa sapidissima

A. 14.0 mmTL

B. 22.9 mmTL

C. 21.0 mmTL

D. 31.8 mmTL 194

Brevoortia tyrannus (Latrobe, 1802) Clupeidae Atlantic menhaden

Range: Atlantic coast of North America from the Gulf of St. Lawrence to northern Florida Meristic Characters Habitat: Pelagic, brackish estuaries and bays as well as inner continental shelf; Myomeres: 45–50 rarely far from land Vertebrae: 45–50 Dorsal fin rays: 18–24 Spawning: May spawn during every month of the year, depending on location, Anal fin rays: 18–24 with peaks during spring and fall; in the study area, most larvae occur Pectoral fin rays: 13–19 in the Middle Atlantic Bight during fall into winter, with more limited Pelvic fin rays: 7 occurrences in coastal waters during spring Caudal fin rays: 7–9+10+9+6–7 Eggs: – Pelagic, spherical – Diameter: 1.30–1.95 mm – Chorion: smooth and thin – Yolk: segmented – Oil globules: single, 0.11–0.17 mm in diameter – Perivitelline space: wide – Late embryo has 2 rows of dorsolateral spots from snout to tail Yolk-sac larva Larvae: – Hatching occurs at 2.4–4.5 mm; eyes unpigmented – Body elongate, with straight gut 70–80% TL; anus always posterior to dorsal fin – Flexion occurs at 8–10 mm – Posterior gut has obvious muscle-band striations – Air bladder evident at about 11 mm – No teeth until 20 mm

– Sequence of fin ray formation: C, D, A – 2P – P1 – Pigmentation includes spots along entire dorsal surface of gut and along ventral surface of posterior half of gut in larvae larger than about 5 mm; few melanophores along dorsum of body disappear at 5–6 mm; pigment present on ventral side of notochord tip; dorsal notochord tip pigment only present in small larvae – When spawned offshore, larvae enter estuaries after about 1 month at sizes of 10+ mm – Transformation occurs at about 30 mm (in estuaries) Note: 1. High number of dorsal and anal rays compared to other clupeids 2. Myomeres between posterior dorsal and anterior anal fins decrease from 5 to 1 3. Brevoortia smithi occurs only as far north as Cape Hatteras; 45–47 myomeres; transforms at 20–23 mm

Early juvenile:

F. 32.0 mm

Figures: Adult: Whitehead, 1977; Egg and yolk-sac larva: Kuntz and Radcliffe, 1917 (redrawn); A–D: Mansueti and Hardy, 1967 (redrawn); E–F: Lewis et al., 1972 References: Houde and Swanson, 1975; Ditty et al. 1994; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 195

Brevoortia tyrannus

A. 8.3 mmTL

B. 10.7 mmTL

C. 16.6 mmTL

D. 23.1 mmTL

E. 27.0 mmTL 196

Clupea harengus Linnaeus, 1758 Clupeidae Atlantic herring

Range: Both sides of North Atlantic Ocean; in the western North Atlantic from Greenland and Labrador to Cape Hatteras; separate stocks oc- Meristic Characters cur in Gulf of St. Lawrence, Banquereau Bank, Scotian Shelf, Gulf Myomeres: 52–62 of Maine and Georges Bank Vertebrae: 52–62 Dorsal fin rays: 16–22 Habitat: Pelagic, schooling; mostly offshore and migrating into deeper wa- Anal fin rays: 15–21 ters during winter; some populations move into coastal waters for Pectoral fin rays: 13–21 spawning Pelvic fin rays: 6–10 Caudal fin rays: 10–13+10+9+8–9 Spawning: Mostly in fall, with a peak in Sep; also a spring spawn, peaking in May; occurs in bays and offshore banks, usually over substrates of rock, cobble or gravel. Georges Bank and periphery of Gulf of Maine are important centers; larvae seldom occur south of Hudson Canyon or off Nw Jersey Eggs: – Demersal, adhesive, clumping, off-round – Diameter: 1.0–1.4 mm Yolk-sac larva – Chorion: smooth, transparent and thick – Yolk: segmented – Oil globules: none – Perivitelline space: wide Larvae: – Hatching occurs at 4–10 mmTL; eyes pigmented – Body elongate with long straight gut; anus always posterior to dorsal fin – Preanus length 80%TL – Flexion occurs at 16–17 mm – Posterior gut has obvious muscle-band striations – Air bladder forms at 5–10 mm, but not obvious until 30 mm

– Sequence of fin ray formation: C – D, A – 2P – P1 – Pigmentation includes melanophores along the dorsal surface of the gut anteriorly, ventral surface of the gut posteriorly; larger larvae have only dorsal gut pigment; early larvae have single streak of pigment on midline of isthmus; large melanophores occur near anus, and at base of caudal fin; pigment present on ventral surface of notochord tip, variable on dorsal surface – Transformation occurs at about 30 mm

Note: 1. High myomere count and late forming anal fin rays are unique for NorthAtlantic clupeids 2. Myomeres between posterior dorsal and anterior anal fins decrease from 8–9 to 4 3. See Mallotus villosus for note on similar larva Early Juvenile:

F. 46.0 mmSL

Figures: Adult: Hildebrand, 1963a; Eggs: Berg et al., 1949; Yolk-sac larva, A–B, D: Krevanovsky, 1956 (redrawn); C: Ehrenbaum, 1909 (redrawn); E: Fage, 1920 (redrawn); F: Susan Kaiser (Able and Fahay, 1998) References: Russell, 1976; Jones et al., 1978; Able and Fahay, 1998; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 197

Clupea harengus

A. 8.2 mmTL

B. 20.2 mmTL

C. 29.0 mmTL Pigment increases on head and dorso-lateral region

D. 29.0 mmTL (Head Region)

E. 41.0 mmTL 198

Dorosoma cepedianum (Lesueur, 1818) Clupeidae Gizzard shad

Range: North American fresh water drainages into the Atlantic Ocean and Gulf of Mexico; reaches estuarine waters of Chesapeake and Dela- ware bays and as far north as Hudson River estuary Habitat: Near surface in open, quiet waters, deeper in fall and winter; found Meristic Characters in submerged vegetation and over mud, sand and gravel substrates Myomeres: 47–51 Vertebrae: 47–51 Spawning: In fresh water in sloughs, ponds, lakes and larger rivers; spring Dorsal fin rays: 10–15 through summer, with peak Apr–Jun Anal fin rays: 25–37 Pectoral fin rays: 14–17 Eggs: – Demersal, adhesive Pelvic fin rays: 7–10 – Diameter: 0.75 mm Caudal fin rays: 9–11+10+9+7 – Chorion: smooth, tough – Yolk: finely granular – Oil globule: 1 large and 1–5 smaller ones – Perivitelline space: narrow Larvae: – Hatching occurs at about 3.2 mm – Body elongate with long straight gut; anus always posterior to dorsal fin – Remnant of preanal finfold still present at 17 mm Yolk-sac larva – Flexion occurs between 11 and 17 mm

– Sequence of fin ray formation: D – C – A, P2 – P1 – Pigmentation includes large melanophores near anus, and a series of spots on dorsal surface of anterior gut; larger larvae develop scattered pigment on caudal fin along base of anal fin, and dorsolaterally on body – Transformation occurs at sizes larger than 24 mm; body begins to deepen at about 35 mm

Early Juvenile: – Scales complete at about 50 mm – Large melanophore forms behind opercle in juveniles – Ventral scutes form at about 35 mm – Note high anal fin ray count

F. 50.0 mmTL

Figures: Adult: H.L. Todd (Goode, 1884); Egg, yolk-sac larva, and B–C: Warner, 1940; A, D: Lippson and Moran, 1974; E: Fowler, 1935; F: Fowler, 1945 References: Lippson and Moran, 1974; Jones et al., 1978 Early Stages of Fishes in the Western North Atlantic Ocean 199

Dorosoma cepedianum

A. 11.0 mmTL

B. 17.5 mmTL

C. 22.0 mmTL

D. 24.2 mmTL

E. 35 mmTL 200

Dorosoma petenense (Günther, 1867) Clupeidae Threadfin shad

Range: Ohio River drainage of North America to fresh waters of northern Guate- mala and Belize; introduced into Chesapeake Bay drainages and occurs in Chesapeake and Delaware Canal Meristic Characters Habitat: Pelagic, schooling, in large bays, lakes, reservoirs, estuaries; in salinities Myomeres: 43–44 0–32 ppt, but mostly below 5 ppt Vertebrae: 43–44 Dorsal fin rays: 11–15 Spawning: Eggs and larvae may be restricted to fresh water, although juveniles Anal fin rays: 17–27 and adults are occasionally collected in estuarine conditions Pectoral fin rays: 12–17 Eggs: – Demersal, adhesive Pelvic fin rays: (7) 8 – Diameter: 0.75 mm Caudal fin rays: 10+9 (PrC) – No other data

Larvae: – Hatching occurs at 4.1–4.4 mm – Body elongate with long, straight gut; anus always posterior to dorsal fin – Flexion occurs at about 10 mmTL Yolk-sac larva – About 36 preanal myomeres

– Sequence of fin ray formation: C, D – A – 2P – P1 (ossification sequence based on illustrations) – Adult fin ray complement present by 18–20 mm – Pigmentation includes a series of melanophores on dorsal surface of anterior gut, and scattered pigment over the anal fin base and caudal peduncle – Transformation begins at about 20 mm

Figures: Adult: Miller, 1963; Yolk-sac larva and A–E: Taber, 1969 References: Lippson and Moran, 1974; Jones et al., 1978 Early Stages of Fishes in the Western North Atlantic Ocean 201

Dorosoma petenense

A. 8.2 mmTL

B. 11.0 mmTL

C. 15.6 mmTL

D. 17.4 mmTL

E. 20.3 mmTL 202

Etrumeus teres (DeKay, 1842) Clupeidae Round herring

Range: Western North Atlantic Ocean from Bay of Fundy to Florida and Gulf of Mexico; also eastern Pacific and Indo-Pacific oceans Meristic Characters Myomeres: 46–50 Habitat: Pelagic, schooling in continental shelf waters; occasionally in bays Vertebrae: 15–17+32–34 and coastal waters, but mostly over outer continental shelf depths Dorsal fin rays: 15–22 Spawning: Not well-described; possibly restricted to south of Cape Hatteras and Anal fin rays: 10–13 Gulf of Mexico; larvae occasionally collected in study area Pectoral fin rays: 14–16 Pelvic fin rays: 8 Eggs: – Pelagic, spherical Caudal fin rays: 10–13+10+9+8–9 – Diameter: 1.17–1.37 mm – Chorion: smooth, transparent and thick – Yolk: segmented – Oil globules: none – Perivitelline space: narrow Larvae: – Hatching occurs at 3.8–4.8 mm; eyes unpigmented – Body elongate with long, straight gut; anus always posterior to dorsal fin Yolk-s ac Larva – Teeth apparent at about 6 mmTL – Snout long and pointed in small larvae – Flexion occurs at 8–10 mmSL – Posterior gut has obvious muscle-band striations

– Sequence of fin ray formation: C, D – A – 2P – P1 – Fin rays not complete until transformation; pelvic forms late and migrates posteriorly – Pigmentation includes about 8 widely spaced, large spots over anterior gut; small spots in double row form over posterior gut; a large melanophore over anus; spot at tip of lower jaw in some larvae – Diagonal streaks of pigment over lower caudal fin base are distinctive – Transformation occurs at 30–33 mmTL Note: 1. Early teeth formation and low anal fin ray count are unique among clupeid larvae 2. Myomeres between posterior dorsal and anterior anal fins decrease from 5 to 2 Early Juvenile:

F. 42.5 mmTL

Figures: Adult: Whitehead, 1977; Egg: O'Toole and King, 1974; Yolk-sac larva: Mito, 1961a; A: Mito, 1961; B: Houde and Fore, 1973; C: Uchida et al. 1958; D: Nancy Arthur (Moser et al. 1996); E–F: Hildebrand, 1963a (Yolk-sac larva and A–C redrawn) References: E.D. Houde, 1981 (pers. comm.); Shaw and Drullinger, 1990; Munroe, 2000; 2002b Early Stages of Fishes in the Western North Atlantic Ocean 203

Etrumeus teres

A. 5.7 mmTL

B. 8.5 mmTL

C. 15.3 mmTL Eye oval in early stages, becomes round at transformation

D. 17.7 mmTL

E. 27.5 mmTL 204

Harengula jaguana Poey, 1865 Clupeidae Scaled sardine

Range: Western North Atlantic Ocean from New Jersey to southern Brazil, including Gulf of Mexico and Caribbean Sea Meristic Characters Habitat: Pelagic and demersal in coastal waters, over mud or sand substrates; Myomeres: 39–42 often in bays, estuaries or high salinity lagoons Vertebrae: 41–43 Spawning: Jan–Sep (peak Apr–Aug); most activity over inner continental shelf Dorsal fin rays: 17–18 (19) at night Anal fin rays: 17–18 Pectoral fin rays: (13) 14–15 Eggs: – Pelagic, spherical Pelvic fin rays: (7) 8 – Diameter: 1.55–1.85 mm Caudal fin rays: 8–9+10+9+7 – Chorion: transparent – Yolk: segmented – Oil globule: single, 0.07–0.10 mm in diameter – Perivitelline space: wide Larvae: – Hatch at about 2.5 mmTL, eyes unpigmented, mouth parts unformed – Body elongate, with straight gut 85–90% TL (decreases to 75% at transformation) – Anus always posterior to dorsal fin – Preanus myomeres 35 at 6.0 mm, 27 at 22 mm – Predorsal myomeres 25 at 6.0 mm, 10 at 22 mm – Flexion occurs at 9–11 mm – Posterior gut has obvious muscle-band striations – Air bladder evident at 6.0 mm

– Sequence of fin ray formation: C, D – A – 2P – P1 – Dorsal fin rays complete at 14–16 mm; anal fin rays complete at 13–15 mm – Pigment sparse on head; early larvae have spots on dorsal (and ventral) tip of notochord, spots at cleithral symphysis, 2 spots near anus, ventral row of spots along posterior gut plus row of spots along dorsum of anterior gut; later larvae form row of spots along dorsum of posterior gut, few spots on bases of caudal fin rays; largest larvae add short rows of spots under dorsal fin and from posterior anal fin on venter of caudal peduncle; few spots also on hindbrain and on "cheek" area – Transformation occurs at about 14–22 mm Note: 1. Low total myomere count; myomeres between dorsal and anal fins 5–7

Early Juvenile:

F. 21.3 mmSL

Figures: Adult: Munroe and Nizinski, 2002; Egg: Gorbunova and Zvyagina, 1975; A, C, E–F: Houde et al., 1974 (redrawn); B, D: Ditty et al., 2006 References: Houde et al., 1974; Gorbunova and Zvyagina, 1975; Fahay, 1983; Munroe and Nizinski, 2002 Early Stages of Fishes in the Western North Atlantic Ocean 205

Harengula jaguana

A. 6.0 mmSL

B. 8.6 mmSL

C. 11.5 mmSL

D. 14.4 mmSL

E. 16.1 mmSL 206

Opisthonema oglinum (Lesueur, 1817) Clupeidae Atlantic thread herring

Range: Western North Atlantic Ocean from southern New England and Bermuda, through the Gulf of Mexico, Caribbean Sea and West Indies; mostly tropical and sub-tropical waters; also eastern Pacific Ocean Meristic Characters Myomeres: 45–49 Habitat: Pelagic, schooling in coastal waters, usually in high salinity, more rarely in Vertebrae: 12–13+32–36 bays or estuaries Dorsal fin rays: 17–22 Spawning: May–Jun off North Carolina; larvae rarely collected in study area Anal fin rays: 20–25 Pectoral fin rays: 15–19 Eggs: – Pelagic, spherical Pelvic fin rays: 8–9 – Diameter: 1.08–1.31 mm Caudal fin rays: 9+10+9+6–7 – Chorion: smooth, thin and clear – Yolk: lightly segmented – Oil globule: single; 0.12–0.16 mm in diameter – Perivitelline space: wide – Paired dorsolateral series of melanophores on late embryo

Larvae: – Hatching occurs at >3.0 mm; eyes unpigmented – Body elongate with long, straight gut; anus always posterior to dorsal fin – Preanus length decreases from 86–91% TL before flexion to 75% SL after transformation – Predorsal myomeres decrease from 25 to 15 – Flexion occurs at about 10 mmSL – Posterior gut has obvious muscle-band striations – Air bladder present, but not obvious until transformation

– Sequence of fin ray formation: C – D – A – 2P – P1 – Pigmentation: in early larvae, melanophores on ventral midline under pectoral fins, posterior to anus, a double row along ventral surface of hindgut, and a dorsolateral row on each side of foregut – Internal pigment over posterior gut well-defined; internal spots over air bladder prominent – Transformation occurs at 15–25 mmSL Note: 1. Myomeres between posterior dorsal and anterior anal fins decrease from 8–10 (<16 mm) to 5–7 at transformation 2. Notochord tip pigment restricted to ventral side 3. High anal fin ray count 4. Pigment increases on dorsum of head at about 24 mmSL Early Juvenile:

H. 30.8 mmSL

Figures: Adult: A.S. Green (Munroe, 2002b); Egg and A–H: Richards et al., 1974 References: Richards et al., 1974; Munroe, 2000; 2002b; Munroe and Nizinski, 2002 Early Stages of Fishes in the Western North Atlantic Ocean 207

Opisthonema oglinum

A. 4.0 mmTL

B. 10.7 mmSL

C. 13.7 mmSL

D. 17.1 mmSL

E. 19.7 mmSL

F. 24.4 mmSL G. 24.4 mmSL (Dorsal Head) 208

Sardinella aurita Valenciennes, 1847 Clupeidae Spanish sardine

Range: Eastern and western North Atlantic Ocean and Mediterranean Sea; in the western North Atlantic from Cape Cod and Bermuda to Brazil, Meristic Characters including Gulf of Mexico and Caribbean Sea Myomeres: 43–48 Vertebrae: 45–49 Habitat: Pelagic in inner continental shelf waters Dorsal fin rays: 16–19 Spawning: Year-round with distinct peaks in different areas; fall–winter off Flor- Anal fin rays: 16–20 ida; Nov–Jun (peak Jan–Feb) off Venezuela; mid-Jun–Sep off North Pectoral fin rays: 15–16 America; Sep–Feb in Gulf of Mexico; larvae uncommonly collected in Pelvic fin rays: 7–8 study area during spring Caudal fin rays: 8+10+9+7 Eggs: – Pelagic and spherical – Diameter: 1.03–1.12 (1.25) mm – Chorion: thin, unsculptured, unpigmented – Yolk: vaguely segmented – Oil globule: single, 0.13–0.18 mm in diameter – Perivitelline space: moderately wide Larvae: – Hatching occurs at 2.5 mm, eyes and body unpigmented, yolk-sac elliptical with oil globule posteriorly – Preanal length ranges from 83–89% SL, then decreases to 73% SL at transformation; head length ranges from 14–27% SL, then increases to 29% SL at transformation; body depth ranges from 8–12% SL, then increases to 25% SL at transformation – Dorsal fin migrates forward: predorsal length 60–68% SL until transformation when it decreases to 41% SL – Myomeres between posterior dorsal fin and anterior anal fin decrease from 7–8 to 5 – Flexion occurs at 7.5 to 9.5 mm

– Sequence of fin ray formation: C – D, A – P2 – P1 – Pigmentation: melanophores occur on nape, over brain and in area of cleithral symphysis; row of elongate melanophores along dorsal aspect of foregut; increasing number of spots along dorsal aspect of hindgut, fewer spots along ventral aspect of hindgut; other melanophores occur along dorsal fin base and lateral body after 16 mm; a few internal melanophores over the anteriormost vertebrae; pigment at notochord tip usually only ventral; pigment on caudal fin increases in larger larvae – Transformation occurs at 16–23 mm Note: 1. See Clupeidae comparative table on Clupeiformes Introductory page 2. Lower anal fin ray count than inOpisthonema oglinum Early juvenile: – In early juveniles, last ray of anal fin is longer than other rays, but last ray of dorsal fin is not elongate; no dark spot on body behind opercle

Figures: Adult: Hildebrand, 1963a; A–F: Ditty et al., 1994 References: Fage, 1920; Matsuura, 1972, 1975; Houde and Fore, 1973; E. D. Houde, 1981 (pers. comm.); Munroe and Nizinski, 2002 Early Stages of Fishes in the Western North Atlantic Ocean 209

Sardinella aurita

A. 3.6 mmSL B. 5.5 mmSL

C. 8.1 mmSL

D. 11.8 mmSL

E. 14.0 mmSL

F. 19.0 mmSL 210

Anchoa hepsetus (Linnaeus, 1758) Engraulidae Striped anchovy

Range: Atlantic coasts of North America and South America from Nova Scotia to Uruguay; absent from Florida Keys and most of Caribbean Meristic Characters Sea Myomeres: 41–44 Habitat: Schooling, pelagic, in estuarine and oceanic waters; mostly over in- Vertebrae: 40–44 ner continental shelf; oceanic occurrences not well-described Dorsal fin rays: 13–16 Anal fin rays: 19–23 Spawning: Spring and summer; Apr–Sep off NE United States, with peak May– Pectoral fin rays: 13–15 Jul; commences near Cape Hatteras, later in southern New England Pelvic fin rays: 7 waters; mostly over inner continental shelf; sporadic occurrences Caudal fin rays: 7–9+10+9+7–8 until Sep Eggs: – Pelagic, elliptical – Diameter: long axis 1.20–1.66 mm; short axis 0.70–0.94 mm – Chorion: smooth and transparent – Yolk: segmented – Oil globules: none – Perivitelline space: narrow Larvae: – Hatching occurs at 3.6–4.0 mm; yolk mass tapers posteriorly – Body long and slender; anus under dorsal fin – Mouth large, terminal, extends to under middle of eye; becomes sub-terminal – Flexion occurs between 5 and 10 mm - Gut with muscle-band striations posteriorly

– Sequence of fin ray formation: C, D, A – 2P – P1 – Pigment very light; series of melanophores along dorsal surface of gut and along anal fin base

Note: 1. Amount of overlap between dorsal and anal fins and number of anal fin rays best characters for distinguishing this species from 2 other engraulid species in study area (see Clupeiformes Introduction) Early Juvenile:

E. 34.0 mmTL

Figures: Adult: Whitehead et al., 1988; Egg, A–B: Hildebrand and Cable, 1930; C: Wayne LaRoche (Farooqi et al., 1995); D: Lippson and Moran, 1974; E: Nancy Arthur (Able and Fahay, 1998) (A, B, D redrawn) References: Able and Fahay, 1998; Berrien and Sibunka, 1999; Munroe, 2000; 2002a; Nizinski and Munroe, 2002 Early Stages of Fishes in the Western North Atlantic Ocean 211

Anchoa hepsetus

A. 3.6 mmTL

B. 5.6 mmTL

C. 10.3 mmSL

D. 15.0 mmTL 212

Anchoa mitchilli (Valenciennes, 1848) Engraulidae Bay anchovy

Range: Atlantic coast of North America from Maine to Florida, through Gulf of Mexico to Yucatan Peninsula, Mexico Meristic Characters Habitat: Schooling, pelagic; estuarine and coastal ocean; sandy beaches, open Myomeres: 38–41 bays, muddy coves Vertebrae: 38–44 Spawning: Primarily in water <20 m deep in bays, estuaries, sounds and coastal Dorsal fin rays: 13–17 ocean; begins as early as Apr off Cape Hatteras, expands rapidly north- Anal fin rays: 23–30 ward over inner continental shelf waters during May and June, occurs Pectoral fin rays: 11–12 as far north and east as Long Island to Narragansett Bay, Rhode Island Pelvic fin rays: 7 in July; declines between September and October Caudal fin rays: 9+10+9+7–8 Eggs: – Pelagic, slightly elliptical – Diameter: long axis: 0.84–1.11 mm – Chorion: smooth and transparent – Yolk: segmented – Oil globules: none – Perivitelline space: narrow

Larvae: – Hatching occurs at 1.8–2.7 mm (smaller than other engraulids); yolk tapers posteriorly – Body long and slender, with anus under dorsal fin – Mouth large and terminal, extends to middle of eye; becomes sub–terminal – Flexion occurs at 7–8 mm – Gut with muscle–band striations posteriorly

– Sequence of fin ray formation: C, D, A – 2P – P1 – Pigment very light, but somewhat more dense than in larvae of 2 other engraulids in study area; series of melanophores along dorsal surface of gut and along anal fin base – Transformation occurs at about 20mm Note: 1. Amount of overlap between dorsal and anal fins and number of anal fin rays best characters for distinguishing this species from 2 other engraulid species in study area (see Clupeiformes Introduction) Early Juvenile:

F. 33.0 mmSL

Figures: Adult: Whitehead et al. 1988; Egg, A, C: Kuntz, 1915; B: Lippson and Moran, 1974; D: Mansueti and Hardy, 1967; E: Fowler, 1945; F: Susan Kaiser (Able and Fahay, 1998) (A–E redrawn) References: E. D. Houde, 1982 (pers. commun.); Vouglitois, et al., 1987; Able and Fahay, 1998; Berrien and Sibunka, 1999; Munroe, 2000; 2002a Early Stages of Fishes in the Western North Atlantic Ocean 213

Anchoa mitchilli

A. 1.9 mmTL

B. 8.4 mmTL

C. 10.0 mmTL

D. 12.0 mmTL

E. 19.5 mmTL 214

Engraulis eurystole (Swain and Meek, 1885) Engraulidae Silver anchovy

Range: Atlantic coast of North America from Massachusetts to northern Gulf of Mexico; also Venezuela to northern Brazil; larvae occur as Meristic Characters far north as Scotian Shelf Myomeres: 43–45 Vertebrae: 25–27+17–19=43–45 Habitat: Schooling, pelagic; mostly over continental shelf waters Dorsal fin rays: 13–16 Anal fin rays: 15–18 Spawning: Spring into fall; not well-described; larvae rarely collected in estu- Pectoral fin rays: 14–16 aries, or as far offshore as Gulf Stream Pelvic fin rays: 7 Eggs: – Pelagic, elliptical Caudal fin rays: ?+10+9+? – Diameter: long axis: 1.02–1.25 mm; short axis: 0.50–0.80 mm – Chorion: smooth and transparent – Yolk: segmented – Oil globules: none – Perivitelline space: narrow

Larvae: – Hatching occurs at 2–3 mm; yolk tapers posteriorly – Body long and slender; anus under dorsal fin – Mouth large, terminal; extends to middle of eye; becomes subterminal – Gut with muscle-band striations posteriorly – Air bladder prominent in larger larvae (Fig. D and F)

– Sequence of fin ray formation: C, D, A – 2P – P1 – Pigment very light, increases with development; note ventral series of melanophores along dorsal surface of gut and along anal fin base; row of elongate spots forms along mid–isthmus in larger larvae; larvae near transformation develop pigment on head, along lateral midline, on caudal fin rays and on dorsal peritoneum – Transformation occurs between 20 and 23 mm Note: 1. Amount of overlap between dorsal and anal fins and number of anal fin rays best characters for distinguishing this species from 2 other engraulid species in study area (see Clupeiformes Introduction) Early Juvenile:

F. 27.4 mmSL

Figures: Adult: Whitehead et al., 1988; Egg, A–C: Kuntz and Radcliffe, 1917; E: Lippson and Moran, 1974; D, F: Markle et al., 1980 (A–C, E redrawn) References: Markle et al., 1980; Able and Fahay, 1998; Berrien and Sibunka, 1999; Nizinski and Munroe, 2002 Early Stages of Fishes in the Western North Atlantic Ocean 215

Engraulis eurystole

A. 3.2 mmTL

B. 3.4 mmTL

C. 5.2 mmTL

D. 18.0 mmSL

E. 22.8 mmTL